. . . we trust that readers will understand our concern with alternate ways of seeing the world that render the same facts in different ways." ( Eldredge & Gould, 1972)
This chapter discusses in detail the evolution of interdependence between long-associated organisms. I present a model that describes evolution to enhancing perpetuity and evolution ensuring compatibility as two different projections of the same complex - a single indivisible process in operation.
In nature there is evidence of co-operation and interdependence. There is a continual coevolution of associated interactors. Two components contribute to the coevolutionary mechanism, as inseparable parts of a single concept such as are the two sides of a single coin in space. PERPETUITY is the easily recognised side of this concept. This is the continual and natural drive or impulse for survival and the perpetuation of the individual. Evolutionary processes also lead naturally to interactions and behaviour that provide a degree of compatibility between long-associated organisms. A compatible animal exhibits behaviour reducing its effect upon the habitat upon which it depends for survival. If measured as a relative value, greater efficiency would be found. Natural ecosystems (such as a coral reef habitat) reflect compatibility in the interdependence of associated organisms.
For an animal to be integrated within a complex system (ecosystem),
it needs to adapt in two ways: there is (i) the classic struggle for survival
and (ii) a "struggle" to be "compatible " with the
A good survivor can do better by being compatible with associated
survivors. When nature devotes millions of years to this, some
interesting solutions can arise. Where introduced species interact with
indigenous plant or animal species, we see the crudest form of this
interaction. With long association, coevolution occurs, where
evolutionary change in one follows the
evolutionary changes in the other (Brewer, 1994). They form an evolving
involving many pairs or large numbers of species. To be termed
changes or responses must be found in both the interacting species
(reciprocity). A moth camouflaged to resemble bark on a tree is not a
coevolutionary tree-moth relationship, yet the moth has adapted to an
evolving, biotic component of
its environment. A bird that has to adapt to find this cryptic moth may
through behavioural changes and improved eyesight. Heliconius sp.
butterfly larvae feed only on Passiflora sp. vines. To deter
laying on their leaves, the vines have evolved fake eggs randomly
distributed on the leaves (Gilbert, 1982). This has evolved as most Heliconius
sp. that deposit eggs have cannibalistic larvae, leading to a
very selective behaviour in egg-laying adults. They avoid leaves with
eggs, as these eggs will hatch first.
This chapter investigates the dynamics that drive
ecosystems leading to the evolution of the complex interrelationships
so critical to ecosystem stability. My emphasis is upon the intensity
of interactions between individual organisms. Just as in studying
the body, we cannot attribute the presence and structure of the heart to mere chance, so in ecosystems
the various associations are not the
result of mere chance or random association. There is an evolved
that leads to holism. Close associations have evolved through time
interdependent systems through necessary reciprocity. Some ecologists,
emphasise stochastic processes, oppose this. Worster, (1994) in his
Nature's Economy notes a modern trend away from a unified theory of
This school sees disturbance, caused by the changing climate, as a
creating mosaics of environmental conditions. They discard the
perspective that preoccupies most of the modeling, theoretical and
work. In this chapter, I hope to show that the evolution of
is an inescapable process that is intrinsic to ecosystems.
, a German biologist (1834-1919), published a book in 1866 describing
how each individual living organism is the product of the interaction
(co-operation) between the environment and the body it has inherited.
He introduced the term, 'oecology'. Ecology as a science
however, did not begin until around 1900 (Allaby, 1986). Ecology has
become the scientific study of the relationships that living creatures
have to each other and to
their environment. Today we
recognise ecosystems, a term introduced by A.E. Tansley in 1935, as
functional units within which animals interact with their physical
environment and each other (Smith, 1990), (Putman, 1994). Ecologists
study communities, populations and organisms in nature, artificial
wheat fields, grain stores, nature reserves), and the consequences of
influence on nature (pollution, global warming) (Begon et al, 1986). In
some modern texts such as Begon, Harper and Townsend's "Ecology,
populations and communities" (1990), they have substituted the
concept with the term community. However, as the colloquial
of a community is something like a group of people or animals living
or sharing something in common, with no emphasis upon the physical
I retain the ecosystem concept. Also, communities do not
necessarily coevolve, while coevolution is an inherent part of an
ecosystem. An ecosystem then consists, as Brewer says
in his book, "The Science of ecology" (1994), "of the community plus
habitat ." A community of organisms is an assemblage of
species and the various interrelationships that bind them. Communities
different populations or groups of species living in the same
Any person living with and depending upon nature is in essence an ecologist. This requires knowledge of edible or poisonous plants, and the behaviour of the animals that they need to catch or evade. New Guinea material culture is 'primitive' (that is stone age) by modern standards, yet a tribesman knows nature intimately, having separate names for about 1,000 different species of plants and animals. They know about animal and plant distribution, and life history (Diamond, 1991). Further, they will have a feel for nature, such as when it is going to rain, the behaviour of various animals and how to live with nature. 99% of our history as humans has been spent as hunter-gatherers, while our technological and industrial society is a very new phase in our history. Urban dwellers are largely detached from the natural environment and have often forgotten the dependence we have upon nature. We need to become ecologists again and learn to understand and live with nature.
Today, just over 100 years after Ecology as a formal science arose, ecosystems are largely disrupted over much of the earth. Biodiversity is declining rapidly. In the chaos discernible, it is difficult to observe the interactive process of perpetuity and compatibility, which requires the absence of human influence. During the interim humanity scrambled to adjust to the new worldview provided by the advances of science, blinded by the acceptance of the idea of survival of the fittest. We have never asked what "fitness" really means. The World Commission on Environment and Development (WCED) expresses a similar sentiment: "When the century began, neither human members nor technology had the power radically to alter planetary systems. As the century closes, not only do vastly increased human numbers and their activities have that power, but major, unintended changes are occurring in the atmosphere, in soils, in waters, among plants and animals, and in the relationships among all of these. The rate of change is outstripping the ability of scientific disciplines and our current capabilities to access and advise" (WCED 1987). One result is that we missed the mechanism of perpetuity and compatibility ; [ glossary ]! An understanding of this mechanism is a tool that we can use to realign ourselves with natural processes.
If you study the living world, you will come to realize an essential characteristic of living organisms is that they live bound into some form of interdependence (Putman, 1994). As Lewis Thomas has observed, "We do not have solitary beings. EVERY CREATURE IS, IN SOME SENSE, CONNECTED TO AND DEPENDENT ON THE REST." Associated animals form ecosystems within which there is the exchange of matter and energy in continuing cycles (Capra, 1982). Nature forms stable associations of organisms that we call ecosystems. These have evolved together over millions of years. Evolution occurs in an ecological context. Evolutionary biologists are interested in the cause of natural mechanisms (McFarland, 1993). Such processes involve behaviour and behaviour often involves social systems or sociality. In its functional, evolutionary context, sociality means more than being gregarious, for it involves the conventions, rituals and relationships of interacting individuals (Wynne-Edwards, 1986). Food chains linking organisms, bind each into this interdependent community or ecosystem. Green plants in their diversity, form the bottom of the food chain. Plants, seaweed or microscopic algae, photosynthesize and so are called primary producers. They use light energy and inorganic compounds to build their tissues. Using the sun's radiant energy, carbon dioxide from the air, and water and nutrients from the soil, plants produce energy-rich sugars. A by-product of the process is oxygen, which plants release into the atmosphere. Plants and animals all need oxygen to survive. Plants synthesise the high-energy organic compounds of their bodies through photosynthesis, so we call them autotrophic ('self-feeding') organisms. Herbivores such as African antelope, Australian kangaroos, American bison, rabbits, caterpillars, some fish and zooplankton all feed exclusively on plants. At this level, we call them primary consumers. Secondary consumers such as birds, lions and fish predators prey upon them. Other essential components to this association are the decomposers, bacteria, fungi and protozoa that break down dead organic matter from plants and animals and so recycle important nutrients back into the living cycle. Primary and secondary consumers breathe out carbon dioxide during their normal metabolism, making this gas again available to plants (Goldsmith et al, 1990).
Ecological science has an important role to play in the study of evolution. E.O. Wilson, a well-known ecologist and author stated, "What we understand best about evolution is mostly genetic , and what we understand least is mostly ecological. I will go further and suggest that the major remaining questions of evolutionary biology are ecological rather than genetic in content. They have to do with selection pressures from the environment as revealed by the histories of particular lineages, not with genetic mechanisms of the most general nature." "I think the greatest advances in evolutionary biology will be made in ecology, explaining more fully in time why the diversity of life is of such and such a nature and not some other." ( Wilson , 1992). Here, he has shown an area of dissension in the evolutionary debate. Niles Eldredge clearly discusses this topic in his book "Reinventing Darwin" (1995).
Ecology as a science is presently in a state of transformation. Among the areas of change are the following:
 The traditional belief that ecosystem complexity begets stability has foundered on theoretical grounds. John Harte: "Many ecologists are disillusioned by the new (mathematical) results discrediting the traditional notion correlating the biotic diversity and the stability of an ecosystem. Yet no current model of which I am aware demonstrates such a correlation; complexity does not promote stability. Complex ecosystems do resist invasions more effectively than simpler systems, since invaders must attack a greater range of organisms, but diverse systems are also given to greater fluctuations, since their stability depends upon a much longer chain of interactions, disruptable at many more junctures" (John Harte, theoretical ecologist, Energy and Resources Group, University of California, Berkeley. (In: Joseph, 1992)). Brewer (1994) offers a clue to the solution to this problem when he says, "real ecosystems . . . because of evolutionary processes, may not correspond to expectations for "systems in general"."
 Competition formed the core of much theoretical development and the question of its validity has caused much disagreement between ecologists and other biologists (Brown & Wilson, 1956). Competition is central to the Darwinian theory of evolution, with "a struggle for existence", and the "survival of the fittest". (Colinvaux, 1973). The debate on competition was central to ecology during the 1970's and 1980's (Begon et al, 1990). Joseph (1990) noted that competition and the evolution of symbiosis are somehow interlinked, for, "to compete, one has no choice but to cooperate and share the pie." One outcome of this debate is a call for "a pluralistic ("more than one" solution) ecology in both theory and method" (Cooper, 1993). Another is the shift towards the acceptance that other influences may be involved, termed non-equilibrium and stochastic (probabilistic) factors. Such influences include physical disturbance and inconstancy in conditions (Begon et al, 1990).
Competition is an anthropomorphic idea. Even in its definition there is some variation in the meanings attributed to it. Brown and Wilson (1956) describe competition as the common striving for some life requisite, such as food, space or shelter, by two or more individuals, populations or species. Simply, they attribute competition to those instances where an organism, be it animal or plant, is seeking, to gain what another is also trying to gain. Smith (1974) adds to this that they interact in a way that affects their growth and survival. Botanists say (check this with plant ecologists) that plants compete for resources, yet in no way does a competitive drive inhere within the plant. The truly objective word is that the plants interact (to their benefit or detriment or with no effect).
Through evolution there is adaptation to the regularities of the environment, be it the abiotic (nonliving physical) environment or the biotic component. Wherever two organisms have niche resources, such as space or food, that overlap, ecologists say they compete. In many ways this is a convenient term, but as it derives from human behaviour, we should not apply it to describe nature. Usually, we can replace the term "compete" by the more objective term, "interact". I will discuss competition in more detail below. In the development of this book, the intensity of interactions has proven to be of crucial importance. In the ecosystem context, this interaction can be with the biotic and abiotic environment. Interactions can also occur between individuals within a population (intraspecific) and between individuals of different populations (interspecific). Intraspecific interactions will prove to be crucial to speciation (species formation).
 Our immense impact upon nature following our rapid overpopulation and technological development has led to visible environmental degradation. With this decline in the quality of our environment has come a need to re-evaluate our relation with nature:
"The question of all questions for humanity, the problem which lies behind all others and is more interesting than any of them is that of the determination of man's place in nature and his relation to the Cosmos. Whence our race came, what sorts of limits are set to our power over nature and to nature's power over us, to what goal we are striving, are the problems which present themselves afresh, with undiminished interest, to every human being born on earth" (T.H. Huxley, 1863) (In: Sagan, 1977).
Important regulatory bodies such as the American EPA (Environmental Protection Agency) have had to refocus their attention and priorities from an emphasis upon "human health" to the goals of monitoring and enforcement activities that protect general " ecological health " as well. America's EPA Science Advisory Board proposed the following policy that aims to realign human activities:
The "EPA should attach as much importance to reducing ecological risk as it does to reducing human health risk. These very close linkages between human health and ecological health should be reflected in national environmental policy. When (the) EPA compares the risks posed by different environmental problems in order to set priorities for Agency action, the risks posed to ecological systems must be an important part of the equation" (Costanza, 1992).
These changes are challenging the human mind to provide a rational and scientific solution. Our need is urgent, for as quickly as we develop new technologies, natural systems are deteriorating into unsustainable states through a loss of ecological integrity and the extinction of species.
 Niles Eldredge, a palaeontologist has identified two camps in the debate of the evolutionary significance of organisation above the population level. I resolve the conflict through providing a mechanism for both speciation (species formation) and the reason for the unchanging persistence (stasis) of some species over long periods of time.
Briefly, the conflict is as follows:
On one side of Eldredge's "High Table" is whom he calls the ultra-Darwinians. They link natural selection intimately with reproductive activities that promote the perpetuation of the individual through the maximisation of its genes relative to other individuals in the same population. They define fitness in terms of reproductive success, with economic success serving this purpose. An organism's fitness is then reflected in the number of genes transmitted to the next generation. This process is defined in terms of active competition between interactors. The logic of this approach is quite clear: no matter how economical, efficient or adapted an organism is, if it does not transmit its genes to the next generation through reproduction, its genetic constitution becomes extinct. Reproduction must therefore define and rule all life's activities. Therefore, competitive activities between individuals take place at the level of the whole organism within the population. All levels, such as ecosystems, above the population level are shaped by this instinct for reproductive success. Dawkins (1982) carries this principle further and claims that organisms are mere vehicles and that the true replicators competing for reproductive success are the genes. To the ultra-Darwinian, evolutionary change is gradual, but constant, as a response to environmental change. Eldredge succinctly describes this as the "traditional (Darwinian) depiction of gradual adaptive transformation" or "phyletic (lineage) gradualism." In this model, speciation (species formation) is a gradual process of slow, steady change by degrees (i.e. phyletic gradualism).
Eldredge defines his opposing view as the "naturalist camp", with natural selection as "a passive ledger recording which variations worked best in the preceding generation." He sees large-scale systems, species, social systems and ecosystems "as absolutely crucial to understanding how the evolutionary process actually works." As a palaeontologist, he finds the patterns in the fossil record that reflect evolutionary histories are not explained by the ultra-Darwinian genetic and population models. As a result of his tracing of fossil lineages, he introduced the idea of "punctuated equilibria" as opposed to phyletic gradualism. He repeatedly found evidence that once species appear in the fossil record, they do not then change much, a phenomenon he termed stasis. He insisted the geneticists needed to provide a model for evolution as it was found to occur in the fossil record, a very valid requirement.
I discuss stasis later in
this chapter, but first I need to introduce
the modified energetic Lotka Volterra model that resolves the conflict
through a new interpretation of the effects of natural selection upon
populations. The model provides an explanation for stasis, and
"Cooperation in the more normal sense has remained clouded by
certain difficulties, particularly those concerning initiation of
from a previously asocial state and its stable maintenance once
A formal theory of cooperation (co-operation) is increasingly needed"
This chapter develops a theory of cooperation that I have
called HOLISM through "perpetuity and compatibility".
In the dimension LIFE, holism is like a coin with two sides,
perpetuity and compatibility, inseparably bound to make a single
reality, mechanism, or process. Darwinian natural selection, is not
unidirectional, but consists of two interlinked processs:
- The individual drive, instinct or impulse to survive,
and perpetuate, generally termed , 'perpetuity', and
- The constraints that result from the dependence and interdependence
of associated and interacting species, termed generally
These constraints result from feedback mechanisms. My theory of PERPETUITY and COMPATIBILITY derives from a study of ecological systems and a study of the science of ecology, especially populations and ecosystems. Ecology requires the scientific study of the INTERACTIONS that cause the distribution and abundance of organisms (Begon et al, 1986). These interactions include environmental influences (abiotic) and other organisms (biotic).
This discussion presupposes the facts of evolution by natural selection , genetic inheritance , the mutability (and therefore variability) of individuals of a species and the reality of ecosystems, communities, habitats and niches in nature. For people of Darwin's time, these were radical or unknown ideas. There was no science of ecology, so the idea I propose had no chance of being found. People believed creation was only 4,000 years old!
Go to next chapter: E. GENETICS and INHERITANCE:
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